Zootaxa 3887 (5): 583–599 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3887.5.5 http://zoobank.org/urn:lsid:zoobank.org:pub:1EA381C9-4FEB-427B-98EA-CBAE8C4362F6 Re-examination of Hemidactylus tenkatei van Lidth de Jeude, 1895: Populations from Timor provide insight into the taxonomy of the H. brookii Gray, 1845 complex (Squamata: Gekkonidae) ANDREW KATHRINER1, MARK O’SHEA2 & HINRICH KAISER3,4 1Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; present address: Depart- ment of Herpetology, Bronx Zoo, 2300 Southern Boulevard Bronx, New York 10460, USA 2Faculty of Science and Engineering, University of Wolverhampton. Wulfruna Street, Wolverhampton, West Midlands WV1 1LY, United Kingdom; and West Midland Safari Park, Bewdley, Worcestershire DY12 1LF, United Kingdom 3Department of Biology, Victor Valley College, 18422 Bear Valley Road, Victorville, California 92395, USA; and Department of Verte- brate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013, USA 4Corresponding author. E-mail [email protected] Abstract Recent herpetofaunal investigations in Timor-Leste revealed populations similar to Hemidactylus brookii Gray, 1845 in four of 13 districts. In order to properly identify these populations, we examined their relationships to other H. brookii- complex populations, notably those from nearby Roti Island, Indonesia (to which the name H. tenkatei van Lidth de Jeude, 1895 has been applied) and topotypic Bornean samples. We evaluated both meristic and mensural data from a set of spec- imens that included the type material of H. brookii and H. tenkatei, and we generated nuclear (RAG1) and mitochondrial (ND2) DNA sequence data for Timor-Leste specimens and a topotypical Bornean specimen presumed to represent H. brookii sensu stricto. Morphologically, Timorese geckos are clearly distinct from H. brookii and identical to H. tenkatei. Our molecular data show that the Bornean specimen thought to be H. brookii is genetically congruent with Timor-Leste specimens, and this specimen is therefore identified as H. tenkatei. Our data also reveal that the Burmese species H. sub- triedroides Annandale, 1905 is distinct from both H. tenkatei and H. brookii. While the current data do not allow us to determine with certainty whether H. tenkatei is the oldest available name for these widespread forms, it is the only name that can be reliably applied at this time. Key words: Hemidactylus tenkatei, H. brookii, H. subtriedroides, Timor-Leste, Borneo, Roti, taxonomy Introduction The genus Hemidactylus Oken, 1817 is one of the most species-rich clades of geckos with a trans-continental distribution (Kluge 1969). Currently represented by 127 species (Uetz 2014), these nocturnal geckos have adapted to a diverse array of habitat types including the tropics and subtropics of Africa, Asia, northern South America, the Caribbean, and Mediterranean Europe (Bauer et al. 2010b). Considering their adaptive plasticity, the majority of Hemidactylus species (approx. 94%) have relatively small distributions in Africa and Asia, while eight species, including H. brookii, H. frenatus Schlegel in Duméril and Bibron, 1836, H. garnotii Duméril and Bibron, 1836, H. mabouia Moreau de Jonnès, 1818, H. persicus Anderson, 1872, H. platyurus (Schneider, 1792), H. parvimaculatus Deraniyagala, 1953, and H. turcicus Linnaeus, 1758, are present in both the New and Old Worlds, having spread throughout these regions through the agency of humans (Carranza & Arnold 2006). Of these, H. brookii has one of the widest distributions and perhaps the most convoluted taxonomic history. Herpetofaunal surveys in Timor-Leste, Asia’s newest country, revealed several brookii-like populations (Fig. 1A) in four of the country’s 13 districts (Kaiser et al. 2011; O'Shea et al. 2012; Sanchez et al. 2012). These were preliminarily identified as H. cf. tenkatei by O'Shea et al. (2012) and Sanchez et al. (2012), given their apparent similarity to H. tenkatei van Lidth de Jeude, 1895, a name coined to identify brookii-complex geckos from the nearby island of Roti. This name had fallen into disuse after being syonymized with H. brookii by de Rooij (1915), but two recent inquiries into the validity of H. tenkatei by Rösler and Glaw (2010) and Mahony (2011) led to the Accepted by L. Grismer: 30 Oct. 2014; published: 27 Nov. 2014 583 FIGURE 1. Specimens of Hemidactylus tenkatei in life. (A) An adult female from Lautém District, Timor-Leste (USNM 579443). Photograph by Mark O’Shea. (B) An adult male from Penang Island, Malaysia (LSUHC 6786), initially identified as H. brookii. Photograph by L. Lee Grismer. (C) An adult male from Sarawak, Borneo (ZRC 2.6167), initially identified as H. brookii. Photograph by Indraneil Das. Images were manipulated in order to show the geckos in similar aspect and similar size. 584 · Zootaxa 3887 (5) © 2014 Magnolia Press KATHRINER ET AL. resurrection of H. tenkatei from the synonymy of H. brookii. Rösler and Glaw (2010) considered H. tenkatei a morphologically distinct species, and Mahony (2011) placed H. subtriedroides Annandale, 1905 into its synonymy. However, during neither of these investigations did the authors examine the three syntypes of H. tenkatei (originally accessioned under the single lot number 4353 in the collection of the Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands). This omission, together with the uncertainty over the identity of populations in Timor-Leste, prompted us to conduct a detailed study of the situation using meristic and mensural characters, as well as a molecular approach. The Hemidactylus brookii complex. Initially described by John Edward Gray in 1845 from specimens ostensibly collected in Borneo and Australia, H. brookii has been considered a species complex distributed across the globe (Carranza & Arnold 2006). Following Gray’s initial description, Murray (1884a, b) described two brookii-like species from what is now Pakistan, H. gleadowi Murray, 1884 and H. kushmorensis Murray, 1884. The first revision of the genus was undertaken by Boulenger (1885), who synonymized multiple African species, including H. verruculatus (Cuvier, 1829), H. cyanodactylus Girard, 1858, H. affinis Steindachner, 1870, and H. guineensis Peters, 1868, with H. brookii, and who placed some specimens identified as H. maculatus Duméril and Bibron, 1836 from a geographically widespread area (India, Sri Lanka, Burma, China) into synoymy with H. gleadowi. Additionally, Boulenger (1890) synonymized H. murrayi Gleadow, 1887 and H. kushmorensis, with H. gleadowi. Boulenger (1898) subsequently synonymized H. gleadowi with H. brookii because he was unable to find definitive morphological characters delimiting H. gleadowi. Through Boulenger’s work, a group divided into at least six species at one time became subsumed under the name H. brookii. Further morphological analyses conducted during the last hundred years have produced multiple conclusions for the origin, dispersal, and differentiation of H. brookii, and they include the descriptions of H. tenkatei by van Lidth de Jeude (1895) and H. subtriedroides by Annandale (1905). Kluge (1969) examined populations on Cuba, Hispaniola, and Puerto Rico and noticed that they differ considerably from their most similar presumptive ancestor from the west coast of Africa; he referred to these conspecifics as a unique subspecies, H. b. haitianus Meerwarth, 1901. These New World “brookii” have experienced their own complex history, but Carranza and Arnold (2006) and Weiss and Hedges (2007) demonstrated that they are members of a deeply divergent lineage of West African H. angulatus Hallowell, 1854. Hemidactylus tenkatei van Lidth de Jeude, 1895. Hemidactylus tenkatei was initially described in 1895 based on a type series consisting of two adults (Fig. 2) and one juvenile from “Rotti” (van Lidth de Jeude 1895). Twenty years later, de Rooij (1915) placed H. tenkatei in the synonymy of H. brookii, but without providing any rationale or comment regarding this change in taxonomy. Hemidactylus tenkatei remained in this synonymy for nearly a century, until Rösler and Glaw (2010) evaluated the morphological variation and taxonomy of H. brookii, H. angulatus, and phenotypically similar taxa based on the holdings of the Zoologische Staatssammlung in Munich, Germany, but without the benefit of an examination of the type specimens of either H. brookii or H. tenkatei. Specifically, Rösler and Glaw (2010) compared Nepalese specimens of what they considered to be H. b. brookii with a male and a female specimen of H. tenkatei from Kupang, West Timor. The morphological comparisons, particularly the differences in three scale characters, led these authors to preliminarily consider H. tenkatei as a valid species, pending further and more detailed examinations. They also stated that their Nepalese specimens were morphologically different from Bornean H. brookii, Borneo being the type locality of the species, ultimately leaving the taxonomic status of the Nepalese population unresolved. Mahony (2011) approached his evaluation of brookii-complex geckos by re-examining the type series of H. brookii and its Asian synonyms. This author provided a description of one “H. brookii” specimen (BMNH 1926.10.30.46) from West Timor, the closest part of Timor to Roti, and considered it indistinguishable from H. subtriedroides, a junior subjective